Gonatodes albogularis (2): NICARAGUA (MNRJ 3164, 10670).
Gonatodes annularis (22): BRAZIL: Amapá: Mazagão, Cachoeira Inajá, Rio Camaipi (MPEG 2667), Oiapoque, km 90 of BR-156, Aldeia Tukai (MPEG 24439), Serra do Navio (MPEG 15080, 15087, 15100, 15148, 19592, 19627-31), Serra do Navio, Área Urucum (MPEG 19213);
Pará: Cachoeira Porteira, Trombetas River (MPEG 16264), Oriximiná, Cruz Alta, 6 km from
Trombetas River (MPEG 15396), Oriximiná, Porto Trombetas (MPEG 24198); FRENCH
GUIANA: Petit Saut, River Sinnamary (MPEG 15826, 15830, 15838, 15841-42); SURINAME:
(MPEG 21823).
Gonatodes caudiscutatus (19): ECUADOR: Guayas: Guayaquil (MZUSP 9181-90, 54370-72), Balzar (MZUSP 9191); Los Rios: Palenque, Rio Palenque Science Center (RMNH 40149-153). Gonatodes ceciliae (5): TRININDAD: Arima, Spring Hill Estate Arima Vallei (RMNH 10160), Maracas, River ly Maracas (RMNH 14913); VENEZUELA: Sucre: San Juan de lãs Galdonas (RMNH 40134-35; MZUSP 53553).
Gonatodes eladioi (1): BRAZIL: Pará: Marabá, Salobo, Serra dos Carajás (MPEG 18020). Gonatodes hasemani (22): BRAZIL: Pará: Juruá, Rio Xingu (MZUSP 67240-57); Rondônia: Porto Velho, Área de inundação da UHE de Samuel (MPEG 15569-71), Porto Velho, Rio Yamary, Cachoeira do Samuel, Território do Guaporé (MNRJ 10678).
Gonatodes humeralis (16): BRAZIL: Pará: Piçarra (MNRJ 16107), Tucuruí, Lago da UHE Tucuruí (MPEG 21881-87). PERU: Loreto: Moropón (MZUSP 28280-87).
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Figure Legends
Figure 1. Dorsal color patterns in preservative in Gonatodes concinnatus complex. (A) pattern A
(MZUSP 54655); (B) pattern B (MZUSP 53674, holotype); (C) pattern C (MPEG 25596) and (D) pattern D (MZUSP 49153). Scale bar = 5 mm.
Figure 2. Ventral color pattern in preservative in Gonatodes concinnatus complex. (A) pattern A
(MZUSP 54655); (B) pattern B (MZUSP 53674, holotype); (C) pattern C (MPEG 25596) and (D) pattern D (MZUSP 49153). Scale bar = 5 mm.
Figure 3. Bivariate scattergrams from stepwise discriminant function analysis (A) using scale
counts and (B) measurements. Gray triangle = Pattern A. Black circles = Pattern B. Black squares = Pattern C. Gray circles = Pattern D.
Figure 4. Gonatodes concinnatus (MZUSP 54655). (A) Dorsal and (B) ventral views of head;
ventral views of (C) right hand and (D) right foot; (E) ventral view of tail. Scale bar = 5 mm.
Figure 5. Gonatodes concinnatus (syntypes, BMNH 1946.9.7.10-12). (A) Dorsal, (B) ventral
views. Scale bar = 5 mm.
Figure 6. Geographic distribution of Gonatodes concinnatus complex. Circles = Gonatodes
concinnatus. Star = Gonatodes tapajonicus. Squares = Gonatodes sp. nov. 1. Triangles = Gonatodes sp. nov. 2. Numbers refer to the localities cited in the text.
Figure 7. Gonatodes tapajonicus (paratype, MZUSP 53674). (A) Dorsal and (B) ventral views
of head; ventral views of (C) right hand and (D) right foot; (E) ventral view of tail. Scale bar = 5 mm.
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Figure 8. Gonatodes tapajonicus from type-locality, Cachoeira do Limão, Pará, Brazil. (A)
Adult male (holotype, MZUSP 53676). SVL = 53 mm and (B) Throat color pattern of male (holotype). (Photos by Miguel Rodrigues)
Figure 9. Gonatodes sp. nov. 1 (holotype, MPEG 25596). (A) Dorsal and (B) ventral views of
head; ventral views of (C) right hand and (D) right foot; (E) ventral view of tail; detail of (F) dorsal scales of trunk and (G) tail. Scale bar = 5 mm.
Figure 10. Gonatodes sp. nov. 1 from Anapu, Rio Xingu, Pará, Brazil. (A) Adult male (paratype,
MPEG 25598, SVL = 46.7 mm), (B) female in life (paratype, MPEG 25599, SVL = 39.7 mm) and (C) Throat color pattern of male (paratype, MPEG 25598). (Photos by Pedro Peloso)
Figure 11. Gonatodes sp. nov. 2 (holotype, MZUSP 49153). (A) Dorsal and (B) ventral views of
head; ventral views of (C) right hand and (D) right foot; (E) ventral view of tail; detail of (F) dorsal scales of trunk and (G) tail. Scale bar = 5 mm.
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Tables
Table 1. Comparative data among color patterns.
Patterns Throat Head dorsally Back and limbs
A Beige, immaculate vermiculations Beige without Brown and beige vermiculations
B (in life, vivid yellow with reddish-White with black oblique stripes brown oblique stripes)
Brown and beige vermiculations (in life, vivid yellow and reddish-
brown vermiculations)
Brown and beige vermiculations (in life,
vivid yellow and reddish-brown vermiculations C
White or dark-brown with black and white oblique stripes (in life, orange with orange yellow oblique
stripes)
Dark-brown and beige vermiculations (in life, orange with orange yellow
vermiculations)
Dark-brown and beige vermiculations (in life, orange with orange yellow vermiculations) D Brown, immaculate vermiculations Beige without Brown and beige small vermiculations
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Table 2. Results of stepwise discriminant function analysis of scale counts comparing the four
groups identified on basis of color pattern. 1DF: loadings on the first principal component; 2DF loadings on the second principal component.
F(+ent,-rem) Wilks's Lambda Approx. F-ratio p-value 1DF 2DF
DSL2T 44.679 0.33 44.679 0 0.65 -0.008 PSL4T 36.711 0.122 40.249 0 -0.057 -0.696 IL 13.583 0.075 34.527 0 0.42 -0.441 PSL2F 9.406 0.052 28.557 0 -0.669 -0.2 VLR 4.36 0.043 25.084 0 0.473 0.056 SL 3.939 0.036 21.598 0 0.072 -0.001
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Table 3. Classification matrix and Jackknifed classification matrix of stepwise discriminant
function analysis using scale counts.
Classification Matrix Jackknifed Classification Matrix Pattern A B C D %correct A B C D %correct Pattern A 26 0 0 0 100.0% 24 2 0 0 100.0%
Pattern B 2 5 0 0 71.4% 2 5 0 0 71.4%
Pattern C 0 0 23 0 100.0% 0 0 23 0 100% Pattern D 0 0 0 14 100.0% 1 0 0 13 92.9% Total 28 5 23 14 97.1% 27 7 23 13 92.9%
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Table 4. Results of stepwise discriminant function analysis of measurements comparing the four
groups identified on basis of color pattern. 1DF: loadings on the first principal component; 2DF loadings on the second principal component.
F(+ent,-rem) Wilks's Lambda Approx. F-ratio p-value 1DF 2DF
MSL 33.213 0.398 33.213 0 -0.561 0.198 HW 18.501 0.215 25.068 0 0.214 0.869 RL 14.107 0.129 23.599 0 0.455 0.579 SSL 8.252 0.093 20.13 0 0.63 0.162 RSW 7.684 0.068 19.317 0 -0.525 0.34 IOD 6.796 0.051 17.962 0 0.528 -0.201
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Table 5. Classification matrix and Jackknifed classification matrix of size-free stepwise
discriminant function analysis using measurements.
Classification Matrix Jackknifed Classification Matrix
Pattern A B C D %correct A B C D %correct
A 26 0 0 0 100.0% 26 0 0 0 100.0%
B 1 4 0 3 57.1% 1 4 0 2 57.1%
C 0 1 22 0 95.7% 0 1 22 0 95.7%
D 1 2 0 11 78.6% 1 3 0 10 71.4%
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Table 6. Snout-vent length and body proportions of Gonatodes concinnatus, Gonatodes tapajonicus, Gonatodes sp. nov. 1 and
Gonatodes sp. nov. 2. Abbreviations are listed in Material and Methods. sd=standard deviation; N=number of specimens. Tail length
of specimens with broken or regenerated tail was excluded. Measurement or
body proportion
Gonatodes concinnatus Gonatodes tapajonicus Gonatodessp. nov. 1 Gonatodes sp. nov. 2
Range Mean sd N Range Mean sd N Range Mean sd N Range Mean sd N
SVL 25.9–48.3 41.60 6.70 26 37.8–51.4 46.10 4.80 7 22.3–55.7 45.00 9.50 23 34.1–45.8 40.90 3.10 14 TL/SVL 1.07–1.16 1.16 0.07 8 1.21–1.23 1.22 0.02 2 1.11–1.26 1.20 0.05 8 1.06–1.32 1.19 0.18 2 HL/SVL 0.23–0.27 0.24 0.01 26 0.22–0.25 0.23 0.10 7 0.21–0.28 0.24 0.02 23 0.22–0.25 0.24 0.01 14 HL/HW 1.30–1.56 1.44 0.06 26 1.45–1.58 1.50 0.06 7 1.21–1.56 1.41 0.08 23 1.44–1.58 1.44 0.05 12 HW/HD 1.31–1.66 1.43 0.10 26 1.28–1.44 1.36 0.05 7 1.17–1.54 1.40 0.10 23 1.25–1.63 1.44 0.11 14 ED/HL 0.19–0.24 0.21 0.10 26 0.21–0.25 0.23 0.01 7 0.20–0.27 0.22 0.02 23 0.20–0.23 0.22 0.01 14 IOD/HW 0.25–0.35 0.31 0.02 26 0.34–0.42 0.38 0.03 7 0.22–0.31 0.25 0.02 23 0.30–0.38 0.35 0.02 14 IND/HW 0.20–0.28 2.10 0.30 25 0.21–0.27 0.25 0.02 7 0.14–0.21 0.17 0.01 23 0.23–0.27 0.25 0.01 14 SSW/SSL 0.82–2.34 1.21 0.28 26 0.99–1.40 1.19 0.16 7 0.88–1.44 1.13 0.13 23 0.91–1.61 1.24 0.19 14 RSW/RSW 1.89–2.35 2.15 0.14 26 1.86–2.21 2.02 0.15 7 1.78-3.01 2.02 0.28 23 1.65–2.19 1.93 0.13 14 MSW/MSL 1.02–1.25 1.13 0.07 26 1.07–1.39 1.21 0.11 7 1.17–1.41 1.28 0.07 23 1.20–1.51 1.32 0.09 14 FL/SLV 0.36–0.42 0.40 0.01 26 0.37–0.44 0.40 0.03 7 0.38–0.44 0.41 0.02 23 0.27–0.40 0.38 0.04 14 HLL/SLV 0.47–0.56 0.51 0.01 26 0.45–0.54 0.51 0.04 7 11.7–28.6 22.70 4.50 23 16.7–21.8 19.70 1.40 14
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Table 7. Scale counts of specimens of Gonatodes concinnatus, Gonatodes tapajonicus, Gonatodes sp. nov. 1 and Gonatodes sp. nov.
2. Abbreviations are listed in Material and Methods. sd=standard deviation; N=number of specimens.
Scales counts Gonatodes concinnatus Gonatodes tapajonicus Gonatodes sp. nov. 1 Gonatodes sp. nov. 2
Range Mean sd N Range Mean sd N Range Mean sd N Range Mean sd N
SAM 109–128 120.6 5.3 25 120–126 122.8 2.3 6 118–143 130.0 6.5 21 100–124 113.5 6.1 14 VVR 49–58 55.0 2.3 25 53–59 55.8 2.2 6 52–60 56.2 2.5 21 44–56 50.4 3.6 14 VLR 17–20 18.5 1.0 25 19–20 19.5 0.5 6 18–24 21.2 1.3 21 17–19 17.7 0.7 14 SL 6–7 6.3 0.5 26 6–8 7.0 0.8 7 5–7 6.2 0.5 23 6–7 6.2 0.4 14 IL 5–7 6.0 0.8 26 5–7 6.1 0.7 7 5–8 6.3 0.9 23 4–6 5.6 0.6 14 PR 3–5 3.5 0.8 26 3–5 3.3 0.8 7 2–4 3.0 0.4 23 2–5 3.1 0.6 14 SP 1–3 1.5 0.8 26 1–3 1.3 0.8 7 0–2 1.0 0.4 23 0–3 1.1 0.6 14 SIS 4–6 5.0 0.4 26 5–7 6.1 0.7 7 4–8 6.0 0.8 23 4–7 5.1 0.8 14 PN 3–4 3.1 0.3 26 3–4 3.3 0.5 7 3 3.0 0.0 23 2–3 2.9 0.3 14 LS 9–13 10.7 0.9 26 10–13 11.1 0.9 7 10–13 11.2 0.9 23 9–11 10.0 0.8 14 PM 2–4 2.3 0.6 26 2–3 2.3 0.5 7 2–3 2.0 0.2 23 2–3 2.3 0.5 14 CM1IL 5–8 6.8 0.9 26 6–8 6.7 0.8 7 6–8 7.0 0.8 23 6–10 7.7 1.3 14 SCS 6–10 8.3 1.1 26 7–10 8.1 1.1 7 7–12 9.4 1.4 23 7–10 8.4 1.0 14 SSC 0–4 1.7 1.5 26 2–4 2.9 0.7 7 0–3 1.1 1.2 23 0–3 2.1 1.1 14 PSL2F 5–7 6.0 0.3 26 6 6.0 0.0 7 4–6 5.1 0.4 23 5–6 5.7 0.5 14 DSL2F 8–12 1.2 1.1 26 10–12 10.3 0.8 7 10–13 11.8 0.9 23 7–11 8.9 1.0 14 SL2F 14–18 16.2 1.2 26 16–18 16.3 0.8 7 15–19 16.9 1.1 23 13–17 14.6 1.1 14 LRS2F 3–4 3.1 0.3 26 3–4 3.1 0.4 7 3–4 3.1 0.3 23 2–3 2.9 0.3 14 PSL3F 6–7 6.3 0.5 26 6–7 6.1 0.4 7 5–6 5.4 0.5 23 5–6 5.9 0.3 14 DSL3F 11–14 12.1 0.9 26 12–14 13.1 0.7 7 13–16 14.3 1.0 23 9–13 11.1 1.2 14 SL3F 17–20 18.6 1.1 26 18–20 19.3 0.8 7 18–22 19.7 1.3 23 15–19 17.1 1.2 14 LRS3F 3–4 3.0 0.2 26 3–4 3.1 0.4 7 3–4 3.2 0.4 23 2–3 2.8 0.4 14 PSL4F 7–9 7.7 0.6 26 7–8 7.4 0.5 7 6–8 6.6 0.7 23 6–8 7.0 0.7 14 DSL4F 10–14 12.1 0.9 26 11–14 12.9 1.1 7 12–16 13.9 1.0 23 9–13 10.7 1.1 14 SL4F 17–22 19.7 1.1 26 18–22 20.3 1.3 7 18–23 20.5 1.2 23 16–21 17.7 1.5 14
78 LRS4F 3 3.0 0.0 26 3–4 3.1 0.4 7 3–4 3.2 0.4 23 2–3 2.7 0.5 14 PSL2T 6–7 6.2 0.4 26 5–6 5.9 0.4 7 5–6 5.3 0.4 23 5–6 5.7 0.5 14 DSL2T 8–11 10.0 0.8 26 10–11 10.4 0.5 7 10–14 12.0 1.0 23 7–11 8.6 1.1 14 SL2T 14–18 16.2 1.0 26 15–17 16.3 0.8 7 15–19 17.3 1.1 23 12–17 14.4 1.2 14 LRS2T 3–4 3.0 0.2 26 3–4 3.4 0.5 7 3–4 3.3 0.5 23 3 3.0 0.0 14 PSL3T 6–8 7.0 0.6 26 6–8 7.1 0.9 7 5–7 6.1 0.7 23 6–7 6.1 0.3 14 DSL3T 11–15 12.7 1.0 26 12–15 13.4 1.0 7 13–17 14.8 1.2 23 10–13 11.4 1.2 14 SL3T 18–22 19.7 1.2 26 19–22 20.6 1.0 7 18–23 20.9 1.3 23 16–20 17.5 1.3 14 LRS3T 3–4 3.1 0.3 26 3–4 3.1 0.4 7 3–4 3.2 0.4 23 2–3 2.9 0.3 14 PSL4T 11–15 12.2 1.0 26 11–12 11.6 0.5 7 9–12 10.3 0.7 23 9–11 9.8 0.8 14 DSL4T 11–14 12.5 1.0 26 11–15 13.4 1.5 7 13–16 14.3 1.0 23 9–13 11.1 1.2 14 SL4T 22–27 24.7 1.5 26 23–27 25.0 1.5 7 23–28 24.7 1.5 23 18–24 20.9 1.7 14 LRS4T 3–4 3.2 0.4 26 3–4 3.4 0.5 7 3–4 3.3 0.5 23 2–3 2.8 0.4 14 RSE 4–5 4.1 0.4 15 4 4.0 0.0 2 4–5 4.4 0.5 16 4–5 4.3 0.5 4
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Figure 1. Dorsal color patterns in preservative in Gonatodes concinnatus complex. (A) pattern A
(MZUSP 54655); (B) pattern B (MZUSP 53674, holotype); (C) pattern C (MPEG 25596) and (D) pattern D (MZUSP 49153). Scale bar = 5 mm.
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Figure 2. Ventral color pattern in preservative in Gonatodes concinnatus complex. (A) pattern A
(MZUSP 54655); (B) pattern B (MZUSP 53674, holotype); (C) pattern C (MPEG 25596) and (D) pattern D (MZUSP 49153). Scale bar = 5 mm.
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Figure 3. Bivariate scattergrams from Stepwise Discriminant Function Analysis (A) using scale
counts and (B) measurements. Gray triangle = Pattern A. Black circles = Pattern B. Black squares = Pattern C. Gray circles = Pattern D.
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Figure 4. Gonatodes concinnatus (MZUSP 54655). (A) Dorsal and (B) ventral views of head;
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Figure 5. Gonatodes concinnatus (syntypes, BMNH 1946.9.7.10-12). (A) Dorsal, (B) ventral
84
Figure 6. Geographic distribution of Gonatodes concinnatus complex, except for Venezuela,
from where no specimens were included in the present study. Circles = Gonatodes concinnatus. Star = Gonatodes tapajonicus. Squares = Gonatodes sp. nov. 1. Triangles = Gonatodes sp. nov.2. Numbers refer to the localities cited in the text.
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Figure 7. Gonatodes tapajonicus (paratype, MZUSP 53674). (A) Dorsal and (B) ventral views
of head; ventral views of (C) right hand and (D) right foot; (E) ventral view of tail. Scale bar = 5 mm.
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Figure 8. Gonatodes tapajonicus, from type-locality, Cachoeira do Limão, Pará, Brazil. (A)
Adult male (holotype, MZUSP 53676). SVL = 53 mm and (B) Throat color pattern of male (holotype). (Photos by Miguel T. Rodrigues)
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Figure 9. Gonatodes sp. nov. 1 (holotype, MPEG 25596). (A) Dorsal and (B) ventral views of
head; ventral views of (C) right hand and (D) right foot; (E) ventral view of tail; detail of (F) dorsal scales of trunk and (G) tail. Scale bar = 5 mm.
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FIGURE 10. Gonatodes sp. nov. 1 from Anapu, Rio Xingu, Pará, Brazil. (A) Adult male
(paratype, MPEG 25598, SVL = 46.7 mm), (B) Throat color pattern of male (paratype, MPEG 25598), (C) female in life (paratype, MPEG 25599, SVL = 39.7 mm) and (D) throat color pattern of female (paratype, MPEG 25599) (Photos by Pedro L. V. Peloso).
89
Figure 11. Gonatodes sp. nov. 2 (holotype, MZUSP 49153). (A) Dorsal and (B) ventral views of
head; ventral views of (C) right hand and (D) right foot; (E) ventral view of tail; detail of (F) dorsal scales of trunk and (G) tail. Scale bar = 5 mm.